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COMMENTARY

Musical pleasure and musical COMMENTARY

Louise Goupila and Jean-Julien Aucouturiera,1

In a pharmacological study published in PNAS, Ferreri et al. (1) present evidence that enhancing or inhibiting signaling using levodopa or risperidone modulates the pleasure experienced while listening to . This result is the latest development in an already remarkable series of studies by the groups of Robert Zatorre and Antoni Rodriguez-Fornells on the implication of the in musical emotions. In their seminal 2001 study, Blood and Zatorre (2) used the PET imaging technique to show that episodes of Fig. 1. (A) The corticostriatal model of musical pleasure of Ferreri et al. (1), peak emotional responses to music (or musical linking the auditory and orbitofrontal cortices to the NAcc, and (B) one of the “chills”) were associated with increased blood many possible mechanisms of musical induction discussed in the in the ventral striatum, the amygdala, and other brain , hypothetically linking the auditory thalami, the amygdala, and the regions associated with emotion and reward. In a dorsolateral prefrontal cortex engaged, for example, when experiencing heavy metal (14). As of yet, it is unclear how mechanisms such as these relate to one 2011 follow-up study, Salimpoor et al. (3) then relied another. It is possible (Left) that they operate independently and that the 11 on [ C]raclopride PET—a technique that allows esti- constructs of musical pleasure and musical emotions are not functionally related. mating dopamine release in cerebral tissue—to show It is also possible (Center) that the dopaminergic model A operates at the same that peak emotional during music listening is level as B and constitutes one of many possible first-order inputs to the construction of the integrative emotional experience (C). Finally, it is also associated with the simultaneous release of dopamine possible (Right) that the model of Ferreri et al. (1) constitutes the evaluative in the bilateral dorsal and ventral striatum. With the process by which the outputs of mechanisms such as B are valuated. Clarifying increasing spatial resolution of fMRI techniques, in these relations will be a major objective for future research. 2013 the same team was able to narrow in on a spe- cific dopaminoceptive subregion of the ventral stria- With a cumulated 4,000 citations since 2001 (bib- tum, the (NAcc) (4). Specifically, liometric data based on a search using Google Scholar they found that NAcc activity during music listening is on January 16, 2019), these four studies (2–5) have associated with how much money participants are had a remarkable influence on the neuroscience liter- subsequently willing to pay for the songs that they ature, and on the music neuroscience community in found pleasurable. In a final salvo to establish not only particular, for which the 2001 study can be considered the correlational but also the causal implication of one of the founding acts: Of these citations, 1,770 dopamine in musical pleasure, the authors have turned (44%) include the words “music” or “musical” in their to directly manipulating dopaminergic signaling in titles. While the question of the musical expression of the striatum, first by applying excitatory and inhib- emotion has a long history of investigation, including itory transcranial magnetic over their in PNAS (6), and the 1990s psychophysiological strand participants’ left dorsolateral prefrontal cortex, a region of research had already established that musical plea- known to modulate striatal function (5), and finally, in the sure could activate the autonomic nervous system (7), current study, by administrating pharmaceutical agents the authors’ demonstration of the implication of the able to alter dopamine synaptic availability (1), both of reward system in musical emotions was taken as in- which influenced perceived pleasure, physiological augural proof that these were veridical emotions measures of arousal, and the monetary assigned whose study has full legitimacy to inform the neurobi- to music in the predicted direction. ology of our everyday cognitive, social, and affective

aScience and Technology of Music and Sound, UMR 9912, Institut de Recherche et Coordination Acoustique/Musique, CNRS, Sorbonne Universit ´e, 75004 Paris, France Author contributions: L.G. and J.-J.A. wrote the paper. The authors declare no conflict of . Published under the PNAS license. See companion article10.1073/pnas.1811878116. 1To whom correspondence should be addressed. Email: [email protected].

www.pnas.org/cgi/doi/10.1073/pnas.1900369116 PNAS Latest Articles | 1of3 Downloaded by guest on September 24, 2021 functions (8). Incidentally, this line of work, culminating in the conscious emotional experiences (15, 16). It is unclear whether article by Ferreri et al. (1), has plausibly done more to attract re- the dopaminergic model should be considered one of search funding for the field of music sciences than any other in these mechanisms, that is, one of many possible first-order this community. inputs to the construction of the integrative emotional expe- The evidence of Ferreri et al. (1) provides the latest support rience (Fig. 1, Center), or whether, alternatively, it consti- for a compelling neurobiological model in which musical plea- tutes the evaluative process by which the outputs of such sure arises from the interaction of ancient reward/valuation sys- mechanisms are valuated (Fig. 1, Right). In the former case, tems (striatal–limbic–paralimbic) with more phylogenetically the so-called expectancy mechanism is of particular relevance. advanced perception/predictions systems (temporofrontal). Given the popularity of these results in the literature, it may In a pharmacological study published in PNAS, come as a , however, that this model is not more tightly integrated with other modern cognitive and psychological Ferreri et al. present evidence that enhancing or views on musical emotions, which, while not contradictory, inhibiting dopamine signaling using levodopa or seem to coexist in relative independence. We would do well, risperidone modulates the pleasure experienced collectively, to consider the paper by Ferreri et al. (1) as a press- while listening to music. ingcalltoreconcileandclarifythe theoretical links between these approaches. This influential theory, which postulates that musical emotion/ Oneelephantintheroomis,first,whetherweareinfact pleasure is computed from the violation of temporal or harmonic talking about identical psychological constructs when address- expectations, was originally formulated by the philosopher of music ing (like in ref. 1) musical pleasure or (like in much of the liter- Leonard B. Meyer (17) but resonates with recent suggestions that ature citing it) musical emotions. Previous work by the same music perception is an active process relying on predictive coding authors has described pleasure as “oneparticularaspectof (18). While Ferreri et al. (1) appear agnostic as to what exact cognitive musically elicited affective responses” (9) and found that it is computation serves as input to striatal activity (citing expectations, strongly correlated to the emotional arousal induced by the but also, e.g., associative conditioning and episodic memory), the music (10). However, strikingly, the manipulation of dopamine well-established fact that striatal activity encodes expectations of used here did not the participants’ reported and reward outcomes (19) has led many to consider these results as arousal, but only their reported pleasure and willingness to pay empirical support to Meyer’s theory. Of the 1,770 music cognition for the music. Thus, how the construction of musical pleasure is articles citing the work, 691 (39%) do so to discuss expectation/ex- linked to, or interacts with, the manifold facets of musical emo- pectancy, and 348 (20%) cite it in conjunction with Meyer (17). In fact, tions studied in the music cognition literature (11) remains un- the expectancy theory of musical emotions has received relatively clear (Fig. 1, Left). Is the construction of musical pleasure/ little direct support (20), and when it has it has implicated the orbito- reward similar for the participants of Ferreri et al. (1) who lis- frontal cortex and amygdala, but not the NAcc (21). Thus, the links tened to the happy, upbeat pop music of Spanish singer Vanesa between expectations and musical pleasure remain underspecified Mart´ın, for the sobbing fans of British singer Adele who find at this stage, and more research will be required to understand how solace in the sad, heart-wrenching inflections of the “Someone exactly predictions relate to dopaminergic release during music Like You” tearjerker (12), and for death metal fans who are able listening. Importantly, and in contrast with previous work by the to convert the growling, heavily distorted sounds of Cannibal same authors, Ferreri et al. (1) show here that dopamine not only Corpse into a pleasing experience of power and peacefulness modulates anticipated emotional peaks or “chills” but also a (13)? How is musical pleasure built from vastly different emo- range of less-intense and more continuous pleasurable epi- tional experiences? More generally, is music always pleasur- sodes. This finding suggests that, beyond peak events such as able? Ferreri et al.’s (1) methodology opens up the possibility chills, musical pleasure is a more continuous phenomenon that to answer these questions by studying whether dopamine- may involve a variety of underlying mechanisms. dependent musical pleasure is a ubiquitous aspect of the mu- However, the final question raised by the work of Ferreri et al. (1) sical listening experience. may lead us even further in elucidating what constitutes the core of Another stumbling block in integrating the corticostriatal our drive to engage with music. While the authors cautiously adopt model of musical pleasure with research on musical emotions is the premise that music does “not seem to have any specific survival its status as a mechanism. It is now widely accepted in the advantage,” many authors, dating back at least to Darwin and Rous- community that several neurologically distinct mechanisms seau, have been interested in the potential functions of music, pro- contribute to the induction of emotions by music (11), all of posing for instance that music plays a central role for sexual which involve separate sensory subsystems (allegedly, for the selection, interpersonal coordination, mood regulation, or the defi- Adele and Cannibal Corpse examples above, the registration nition of self-identity (22). The finding that music constitutes a priv- of sad pitch contours in the voice areas of the right superior ileged stimulus able to activate phylogenetically ancient systems temporal gyrus for the former and the rapid activation of the involved in valuation and motivation may very well be interpreted amygdala by thalamically encoded cues of auditory threat for as an indication that the braincontainsanadaptiveneural the latter; Fig. 1). One tentative model is that different as- specialization for processing music as a rewarding stimulus. As such, pects of music, including acoustic signals (e.g., low-pitch and one might whether the crucial question for future research is low-frequency content for Adele and roughness for Cannibal not so much whether music is rewarding, but rather why. Corpse), harmonic and temporal structures (e.g., minor chords and slow pace versus fast pace), familiarity, and so on, are inter- Acknowledgments preted in parallelly working subsystems, before being cogni- This work was supported by European Research Council Grant StG 335536 tively interpreted and integrated to give rise to idiosyncratic, CREAM and Agence Nationale de la Recherche programs MICA and REFLETS.

2of3 | www.pnas.org/cgi/doi/10.1073/pnas.1900369116 Goupil and Aucouturier Downloaded by guest on September 24, 2021 1 Ferreri L, et al. (2019) Dopamine modulates the reward experiences elicited by music. Proc Natl Acad Sci USA 116:3793–3798. 2 Blood AJ, Zatorre RJ (2001) Intensely pleasurable responses to music correlate with activity in brain regions implicated in reward and emotion. Proc Natl Acad Sci USA 98:11818–11823. 3 Salimpoor V, Benovoy M, Larcher K, Dagher A, Zatorre R (2011) Anatomically distinct dopamine release during and experience of peak emotion to music. Nat Neurosci 14:257–226. 4 Salimpoor VN, et al. (2013) Interactions between the nucleus accumbens and auditory cortices predict music reward value. Science 340:216–219. 5 Mas-Herrero E, Dagher A, Zatorre RJ (2018) Modulating musical reward sensitivity up and down with transcranial magnetic stimulation. Nat Hum Behav 2:27–32. 6 Seashore CE (1923) Measurements on the expression of emotion in music. Proc Natl Acad Sci USA 9:323–325. 7 Krumhansl CL (1997) An exploratory study of musical emotions and psychophysiology. Can J Exp Psychol 51:336–353. 8 Kraus N, White-Schwoch T (2017) Neurobiology of everyday communication: What have we learned from music? Neuroscientist 23:287–298. 9 Zatorre RJ, Salimpoor VN (2013) From perception to pleasure: Music and its neural substrates. Proc Natl Acad Sci USA 110:10430–10437. 10 Salimpoor VN, Benovoy M, Longo G, Cooperstock JR, Zatorre RJ (2009) The rewarding aspects of music listening are related to degree of emotional arousal. PLoS One 4:e7487. 11 Juslin PN, Västfjäll D (2008) Emotional responses to music: The need to consider underlying mechanisms. Behav Brain Sci 31:559–575. 12 Vuoskoski JK, Thompson WF, McIlwain D, Eerola T (2012) Who enjoys listening to sad music and why? Music Perception Interdisciplinary J 29:311–317. 13 Thompson WF, Geeves AM, Olsen KN (2018) Who enjoys listening to violent music and why? Psychol Popular Media Culture, 10.1037/ppm0000184. 14 Ollivier R, Goupil L, Liuni M, Aucouturier J (2019) Enjoy the violence: Is appreciation for extreme music the result of cognitive control over the threat response system? bioRxiv:510008. 15 Barrett LF (2017) The theory of constructed emotion: An active inference account of interoception and categorization. Soc Cogn Affective Neurosci 12:1–23. 16 LeDoux JE, Brown R (2017) A higher-order theory of emotional consciousness. Proc Natl Acad Sci USA 114:E2016–E2025. 17 Meyer LB (1956) Emotion and Meaning in Music (Univ of Chicago Press, Chicago). 18 Koelsch S, Vuust P, Friston K (2018) Predictive processes and the peculiar case of music. Trends Cognit Sci 23:63–77. 19 Pessiglione M, Seymour B, Flandin G, Dolan R, Frith C (2006) Dopamine-dependent prediction errors underpin reward-seeking behaviour in . Nature 442:1042–1045. 20 Egermann H, Pearce MT, Wiggins GA, McAdams S (2013) Probabilistic models of expectation violation predict psychophysiological emotional responses to live concert music. Cognit Affective, Behav Neurosci 13:533–553. 21 Lehne M, Rohrmeier M, Koelsch S (2013) Tension-related activity in the and amygdala: An fMRI study with music. Soc Cognit affective Neurosci 9:1515–1523. 22 McDermott J (2008) The evolution of music. Nature 453:287–288.

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